Wednesday, January 17, 2018

The Daylily as a Permaculture Subject - Part 3

The Daylily as a Permaculture Subject 

Part 3
Breeding for Improvement 

Brian Reeder

*Disclaimer - (I am not recommending in any way, form or fashion that anyone should consume any part of the daylily. While Hemerocallis (daylily) is a common food and medical substance in Asian countries, if you have never eaten daylily or do not have accurate information on the consumption of daylily, it is completely upon your discretion when experimenting with consuming daylily. I strongly recommend that first time consumers be very careful in the amount of any given daylily they eat in order to observe the effects on their individual systems. Some people have experienced diarrhea from consuming daylily in quantity and some daylilies have been known to have sedative effects, which might be referred to as ‘poisoning’ or ‘poisonous effects’ by some sensitive people. The internal chemistry of each daylily, as well as each person, may vary and so no general information about daylilies as food or medicine can be made that will cover all daylilies and all human reactions/interactions with/to daylilies in any given instance. What is perfectly fine for one person may not be for another.)

In the final installment of this series we will look at the possibilities of forming a breeding program to select for traits in daylilies that can increase their value to permaculture, both in terms of desirable plant traits and nutraceutical traits.

I have mentioned several times in the past two articles that I have been making some selection in this direction, with a focus on plant traits that make for a better garden as well as permaculture specimen and flower traits that make for better display uses as well as culinary applications. This does not mean that any of these plants are separate from flower trait selection (in terms of floral display) in my overall floral daylily program. The traits that make for a better permaculture specimen also apply to making a better and more garden-worthy floral specimen. As well, one of the most desirable traits in a good culinary daylily flower is also of utmost importance to a good garden specimen of floral display daylilies - i.e., resistance to thrips, aphids and other insect predators that cause damage to or destroy the flowers, buds and sometimes even scapes.

A hybrid daylily with high susceptibility to thrips, showing extreme damage to buds and scapes, and foliage heavily damaged from late freezes. I suspect that the combination of late freeze susceptibility and thrip susceptibility can combine to make a much more severe effect. You can see the beautiful, deep green foliage of the plant in the background to the right, which has strong late-freeze resistance and is a later blooming cultivar and so showed no damage. 

So my permaculture selection isn’t a separate part of my program, but rather, is composed of plants from within my overall selection program that are attractive, vigorous, hardy and show high resistance to disease and insect pests and just happen to also have very good tasting buds/flowers that have no negative effects on me physically when I eat them. So while there may be people out there who’s only interest in daylilies would be to breed for permaculture applications (and may select for a specific type of flower in conjunction with their program, or not), anyone who is breeding daylilies for any reason, who pays any attention to plant traits, could theoretically be selecting plants that also show high culinary value (though I am not suggesting everyone should be selecting for culinary or permaculture traits, either). However, such selection does not have to be a ‘separate thing’ from daylily flower breeding, though it can be. That depends on the interests of the breeder. I just want to stress that this does not have to be a separate thing, a whole world unto itself, unless the breeder wants it to be.

The crosses that have shown the greatest application for culinary uses in my program, and that are also giving me the best range of desired plant and flower traits, are the base crosses I have made to build my future floral daylily program upon. In fact, the plants that are showing the best applications for permaculture purposes in my program happen to be a side effect of the outcrossing I have done between modern hybrids over species and species-like material - specifically seedlings coming from Hemerocallis fulva Korean, Hemerocallis Hankow, Notify Ground Crew and Ancient Elf.
So this is how my own breeding for permacultural uses emerged - by chance as a secondary side effect of my program to build a new tetraploid base-stock by going back to the species and species-like hybrids to open the tetraploid gene pool. The selections I have made are selections I would have been making anyhow, and my focus has been a vigorous, hardy plant with high disease resistance and a lovely flower with high pest resistance. By chance some of those selections have also turned out to be delicious, but if one wanted to make actual crosses and selections toward specific goals of permacultural or nutraceutical uses, that would be possible as well. In fact, I am now beginning to make some crosses of this nature, now that I have plants that I feel would have special application for these purposes.

So what could one be selecting for? On the most basic level, one could be making crosses and selection for the most basic and desirable traits - hardiness, vigor and disease resistance of the plant and flowers of good flavor with high pest resistance. However, there are probably many other traits that could end up being bred and selected for. Since (apparently) no one in the west is actively breeding and selecting for nutraceutical traits, it looks to me as if this would be an open field, where almost any type of program could be ‘unique’ or nearly so.

In terms of food applications, some daylily flowers taste very sweet, almost like very sweet lettuce, while others are more bitter, less sweet or more astringent in flavor. Some say that one color or another is sweet and some others are bitter, etc., but I have not found this to be the case, and I cannot say to you that a given color is “good” while another is not. I find it is much more variable - one yellow is delicious, another is less so, one purple is bitter while another is sweet, etc. The species and near-species hybrids that I have previously discussed all taste good to me, for instance. 

Some daylilies have been shown to have high protein levels and high vitamin C levels in their flowers, but I don’t know of any testing across many types to determine variations. This may be a open field for selection. Other flower traits that might be selected for, combined with flavors and nutritional content, would be flower size, texture and storage ability. A delicious, nutritious and large flower could be very enticing, especially combined with high pest resistance on a vigorous plant with disease resistance, hardiness and many buds on the scapes. Let your imagination be your guide and experiment with what is possible. I expect a great deal is possible once attention is paid to the underlying trait variations.

Whooperee - An exceptionally thrip and rust resistant evergreen cultivar.

In experimenting with eating daylily flowers I have also run across some interesting effects, in addition to flavors. As mentioned in the first part of this series, daylilies have traditionally been attributed with medicinal properties, especially sedative effects, and modern research (as noted in that first part) is also beginning to demonstrate that this may well be the case. In my own experience, this is the case, and the effect(s) and strength varies from one plant to the next. Some plants have little to no effects at all, and these are the most desirable, in my opinion, as a food source. Others will have a mild sedative effect that manifests for me as a slight haziness that is relaxing but does not interfere with my ability to function. 

Still others have a considerably stronger effect and these can be so strong as to be described as strongly sedative and even mildly hallucinogenic, creating a euphoria combined with visual lights and colors, especially when the eyes are closed, and stimulate very deep and relaxing sleep, with no notable effect upon waking the next day. It is my experience that these are quite rare and are probably responsible for the reports of daylilies that are ‘toxic’ or ‘poisonous’. However, I have not experimented with any of these types enough to give any definitive report and I do not recommend you try any daylily flowers except in very small doses (one petal, perhaps?) until you find its effect on you, as well as your body’s tolerance to daylily flowers in general. If you see no negative effects, you might increase the amount you eat at a given time to see if there is a dosage effect in larger quantities.

The strongest effects I have experienced have been from eating five or more buds/flowers (and fresh buds seem to have stronger effects than dried and cooked flower) so perhaps some of the ones with milder effects would show stronger effects in larger quantities. However, since these pharmaceutical effects do not hold much interest for me, I have not experimented very much with these effects and so can give very little information and can offer no recommendations, except to say, that for those who might have an interest in these effects, whether permaculturalists, medicinal/pharmaceutical researchers or interested hobbyists, there is seemingly variation and undoubtedly these are areas that can be modified (perhaps highly so) through selection. Good research in labs by professional researchers would be of much value, both for pharmaceutical effects and nutritional factors. Without access to such testing, hobbyists and permaculturalists can still make progress on selection for such traits, though I strongly recommend they proceed with caution, especially in terms of the pharmaceutical applications.

Further, as the research cited in the first part suggests, there may be even more pharmaceutical effects in Hemerocallis and these traits may well be amenable to selection as well. However, I feel these may be more difficult to select for, unless one has access to laboratory testing. I personally wouldn’t have a clear idea of how to do such selection without access to chemical and perhaps genetic testing, as well as application trials in non-human test subjects. I think this work is beyond the scope of most hobbyists, but may be applicable for some permaculturalists, especially where that person(s) is a trained researcher with access to lab testing. I want to stress that I am not suggesting anyone should attempt to do work with traits of a chemical nature that they are not qualified to work with, but these traits may well exist in the Hemerocallis, and if one has the skill-set to work with such traits, I would still hope they would apply the other breeding suggestions I am recommending herein for working those traits into strong, vigorous and disease/pest resistant lines.

The traits and uses that I want to stress here, both because I think they are the most applicable traits for hobbyists and home permaculturalists to work with and because they are the traits I am personally focused on and feel I can comment on best, are the plant traits that make a great plant for permaculture/food forests/landscape specimens and that make flowers stronger, durable, resistant to pests and useful as a floral or food item. I feel these are easy traits to work with and can have the greatest application. Both are complementary for those already breeding daylilies for flowers and both will be of value to those who solely have an interest in daylilies for permaculture applications. 

(Gram's Dream x Arctic Snow)
A seedling from my program showing the type of plant, scape, bud count and flower I like - strong, hardy, healthy, disease resistant, brightly colored and unblemished.

With that said, I do want to stress that I feel permaculture shouldn’t eschew beauty and can engage in any level of flower phenotype selection that might tickle the fancy of that permaculturalist. I do not know if all daylily phenotypes will have applications for food purposes. I haven’t eaten flowers from every daylily cultivar or seedling I grow. Some flower phenotypes may be pleiotropic with, epistatic to or linked to traits that would cause problems with food applications. However, where any flower trait can be combined with good food qualities, then the sky is the limit.

Fortune's Dearest 
A lovely purple color and white tooth edge to petals and sepals - a lovely look from modern floral daylily breeding.

I also want to stress that I am not preparing this series to create sales for myself of either daylily species or cultivars or my own introductions. My goal is to inspire others to look at and experiment with the daylily beyond those who are already floral daylily hobbyists, and with floral hobbyists, beyond the singular focus on flower appearance. I do sell H. fulva Korean, Notify Ground Crew and Ancient Elf, but my stock is small and I have no interest in becoming a large commercial seller, nor do I have any interest in selling out of these. I do want to promote these plants and I want to see many people growing and breeding from these and other good cultivars, but whether they are sourced from me or another vendor is irrelevant. What is relevant is to spread the word about what I am finding that is working best for me and for other breeders whom I communicate with. I hope to see many other growers and vendors offering these, regardless of where they obtain their initial stock for propagation.

Further, I will be introducing plants in time that I consider highly applicable to permacultural uses. In all instance, where I know the parentage (which will be most of the time), I will list that parentage. In that way, you can buy these cultivars from me (and I do hope to spread my best work around for others to grow and use), but more importantly, I want to get the information out to others as to what I have worked with and what has given me the best results so that others can work within those same genomes and use them to make their own unique combinations and selections. Further, my best selections will not always fit every environment, so others need to be tailoring this type of breeding for their own environments, even if they do work with some of my introductions. 

(H. fulva Korean x Queen's Circle)
A seedling and future introduction from my program that has many applications for both floral and permaculture breeding. 

My goal is to increase knowledge and interest, as well as to inspire others to also pursue work beyond the increasing and intensification of the flower traits. If you want to buy the best results of my breeding, testing and selection, I certainly will not complain, and I understand that some people will not want to take on a breeding and testing program of the scale I have and so may prefer to buy improved stock rather than breed it up themselves. However, for those who wish to do the breeding themselves, take the information I offer and run with it! Take it in directions I have tried and in directions I have not tried. Put your own unique stamp on it and select it to your specific garden needs and your own tastes and aesthetic. 

(Notify Ground Crew x Arctic Snow)
A seedling and possible future introduction from my program that has many applications for both floral and permaculture breeding.

More than anything, I want to inspire people who might not really think much of daylilies to begin thinking about them and growing them. They have many uses beyond flower gardens. Since there is little available information about these other uses, it is my hope to spread some of what I have found through trial and error and to offer you useable information to make good selections. The greatest asset of the hybrid Hemerocallis is that there are a great many varieties and they show great diversity. However, this very thing can also be their greatest drawback, as they are not all created equally and making good selections for uses beyond the flower can be extremely difficult due to the lack of reliable information. I hope to help make that information a little more accessible and to inspire you to seriously consider the daylily.

Now I want to touch on my experiences and offer an assessment on a few of the traits I feel are important to permaculture uses (and floral daylily breeding, as well).

1. Rhizomatous Spreading - Usually said to be recessive in the floral daylily community, but my observation is that the trait appears to be a single major dominant gene with simple Mendelian behavior. There are likely modifiers as well, but the appearance is of a single gene that is dominant with a dosage effect. Homozygotes show greater spreading behavior (thought this may vary in just how much, how far and how vigorously - thus the possibility <likelihood> for modifiers). Heterozygotes show spreading behavior, though less so, spreading slower and each fan arising at a shorter distance from the main plant. All the fulva clones I have grown are rhizomatous to some extent or another. Some daylily hybrid cultivars show rhizomatous behavior such as Bill Fall and La Vida Loca. The trait is generally considered undesirable by floral daylily hobbyists, but can have many applications for landscaping uses and permaculture uses.

A patch of rhizomatous daylilies

Hybrid tetraploid daylily cultivar Bill Fall

H. fulva Korean growing in large containers. This is an excellent method to restrain rambunctious rhizomatous types of daylilies. Some running types of daylilies can escape and become wild, invading surrounding areas, so care should be taken when growing such types if their escape is a concern for you. Containers allow you to maintain such plants without fear of escape. Just keep an eye on any runners that try to emerge from the holes in the container to be sure they don't establish in the surrounding soil. Not all running types are invasive enough to be a concern, with H, fulva ex Europa, commonly called the ditch lily and ubiquitous throughout much of the world, being the most extremely invasive type.

2. Clump Forming - While it is generally thought of as dominant in the floral daylily community, clump forming seems to be recessive to rhizomatous growth, in my experience. All of the yellow species are clump forming and most hybrid daylilies are clump forming. In tame flower gardens, clumping is generally preferred, but in many permaculture applications, the spreading habit of the rhizomatous types may actually work better.

A beautiful display from the clump-forming daylily cultivar Spider Man. Spider Man is a fairly dormant plant that shows high thrip and rust resistance in my experience.

3. Rust Resistance - Seems to involve multiple genes, both dominant and recessive. Daylily plants that show very high rust resistance in field tests can often show very good breeding value for rust resistance. Because of the multigenic nature of rust resistance, I can’t tell you any percentages to expect, as that will vary wildly. The genetics of rust resistance is complex. However, I can recommend that H. fulva Korean and Ancient Elf have shown exceptional rust resistance and breeding value for rust resistance in my own program. To test daylilies for rust resistance, you must expose them to daylily rust and you must not spray them with fungicides to combat rust. To breed for rust resistance, it is best to use the most resistant plants you can find as parents, and their seedlings must be exposed to rust. Without rust, you can’t really select for resistance to rust. However, even if you can’t select for rust resistance, you can at least base your lines on plants that show exceptional rust resistance.

A susceptible plant to daylily rust (behind) showing heavy infestation with a very highly resistant daylily plant (foreground) showing no rust infection at all.

4. Thrip/Pest Resistance - While I have seen daylilies that appear to be fully immune to rust, I have never seen any daylilies that seem to be totally immune to thrips. However, I have seen a few cultivars and seedlings that show high thrip resistance. I have been breeding and selecting for thrip/pest resistance and it seems that those plants that show very high resistance to thrips are able to produce seedlings that also show very high resistance. I cannot yet say that this trait is a single gene or multigenic, or that it is clearly dominant or recessive. What I can say is that it is apparently heritable. I will need to observe for some years more and with a good number more crosses to have a good feel for what to expect in breeding and selecting. What I can tell you now is that the trait(s) appears to be heritable when thrip/pest resistant plants are used as parents. The most thrip resistant plants I have found include Solaris Symmetry, Whooperee, Spider Man, Notify Ground Crew, Tis Midnight and Corky (and its descendants including From Darkness Comes Light, etc.) As with rust resistance, you need to expose daylilies to thrips and other insect pests to select for resistance and you must not spray pesticides to eliminate these pests.

Hybrid Daylily cultivar Solaris Symmetry

5. Flower Flavor - I have only been selecting for this trait for a couple of years and haven’t seen enough seedlings to make a guess as to how many genes may be involved or how those genes may work, but I can say that the trait seems to be heritable. In other words, daylilies that taste good to me have produced seedlings that taste as good or nearly so, though sometimes they also produce daylilies that have flowers that do not taste good. All I can tell you at this time is that there seems to be heritability for the flavor traits. I will address this more in the future as I have a better idea of what is happening, but I wanted to tell you that there is apparently room here for selection, so please, do try if you are interested.

6. Flower Color (Visual Anthocyanin versus Non-Visual Anthocyanin) - I want to address this in the most general of terms. This is actually a very complex area, but at the most basic level, we can look at the two classifications the Chinese use ( ‘yellow’ vs ‘orange’). For me, ‘yellow’ encompasses all the non-visually anthocyanic types such as all tones of yellow (carotene) and all tones of ‘melon’ (a seemingly-recessive gene that changes carotene to lycopene), pigments that are deep within the tissue of the flower petals and are also present in the visible-anthocyanic group in the petals underneath the anthocyanin serving as ‘base-color’ that can effect the visual tones of the anthocyanin as perceived by our eyes. 

The ‘orange’ group includes all those that have visible anthocyanin (a water-soluble pigment in the upper level of the petal) ranging from orange to red to purple to blackish-red to blackish-purple and also including all pink and lavender tones and possibly some near white flowers (near white is likely occurring on both anthocyanic and non-anthocyanic types). I also want to mention that I refer to “non-visual anthocyanin” (the ‘yellow’ category) as such because these almost certainly contain anthocyanin that, as shown by putting these flowers under ultraviolet light, would be visible to insect eyes, but not to human eyes. Visual anthocyanin may be present on the entire flower, only on the petals (and not on the sepals or vice-versa) or as an eye, either on visually anthocyanic types or on non-visibly anthocyanic types.

I am not going into details about the heritability of all these different variations. What I want to focus on here is the most basic level. I suspect that the fulva is older, evolutionarily speaking, than the many yellow species. I suspect that non-visible anthocyanin is a simple ‘knock-out’ gene. That is, a gene that stops the visible anthocyanin from forming in the flower. My experience is that visible anthocyanin is dominant, while non-visible anthocyanin (or absence of visible anthocyanin, if you prefer) is recessive and that this is a simple, Mendelian single gene. The complexity comes after one or the other of these two pathways are active. There are other genes that determine which type of anthocyanin you get (each color of anthocyanin is likely something different genetically and are different types of anthocyanin such as delphinidin, which makes purple tones, etc.), while yet other genes are responsible for eyes, bicolor, self, dilution of tone/intensification of tone, etc. 

An example of a yellow daylily (left), showing no visible anthocyanin, (center) under ultraviolet light where patterning that may be anthocyanic in nature is revealed. Picture right is the same flower in grayscale.

My experience is that at its most basic level you either have visual anthocyanin (which is dominant) or you don’t have visual anthocyanin (which is recessive). This actually makes breeding fairly easy. At this most basic form, let’s do one example - yellow and red (of whatever shade or tone). A red, having visible anthocyanin which is dominant, can have one or two doses (at diploid level) and still be visually red (of whatever shade or tone). When crossed to the yellow flower without visual anthocyanin (no eyes, etc., just yellow self), the homozygous-for-visible-anthocyanin-red creates 100% of its seedlings that show visible anthocyanin (red, orange, reddish tones, red or orange bicolor, red or orange eye, etc., depending on the other modifier genes present). When the yellow is bred to a red that is heterozygous for visible anthocyanin, you will see 50% seedlings showing visible anthocyanin and 50% of seedlings that do not show visible anthocyanin. Some of the 50% showing visible anthocyanin may just be an eye or bicolor, etc., depending on the other genes present. Tones of each may vary, but the split at this basic level seems to be quite reliable across a huge number of crosses I have made.

Crosses between different tones/colors of visibly anthocyanic flowers or between different tones of non-visibly anthocyanic flowers are not simple and can produce a wild range of tones. This indicates to me that there are multiple genes involved in the exact color and tone of visible anthocyanin and the non-visibly anthocyanic, as well. Neither group seems to be straightforward or simple in terms of exact colors, combinations and patterns and must represent multiple genes.

I hope this gives you a basic overview. I have tried to give you some suggestions for base plants, plants for rust resistance, plants for thrip resistance and some idea of what to expect (at the most basic level) in terms of breeding for some of the basic traits that may be of interest in breeding daylilies for permaculture purposes. For anyone who is interested in pursuing this type of daylily use or breeding, please feel free to contact me through my Facebook daylily page, Sun Dragon Daylilies, or through the email address found on the about/contact page at my website Sun Dragon Daylilies. There is much more to the daylily than just the flower and I hope a new generation of breeders will pursue these traits in addition to the beauty of the flower.

Part 1     Part 2

Sunday, January 14, 2018

The Daylily as a Permaculture Subject - Part 2

The Daylily as a Permaculture Subject 

Part 2
Selecting Plants for Growing and Base Plants for Breeding

Brian Reeder

*Disclaimer - (I am not recommending in any way, form or fashion that anyone should consume any part of the daylily. While Hemerocallis (daylily) is a common food and medical substance in Asian countries, if you have never eaten daylily or do not have accurate information on the consumption of daylily, it is completely upon your discretion when experimenting with consuming daylily. I strongly recommend that first time consumers be very careful in the amount of any given daylily they eat in order to observe the effects on their individual systems. Some people have experienced diarrhea from consuming daylily in quantity and some daylilies have been known to have sedative effects, which might be referred to as ‘poisoning’ or ‘poisonous effects’ by some sensitive people. The internal chemistry of each daylily, as well as each person, may vary and so no general information about daylilies as food or medicine can be made that will cover all daylilies and all human reactions/interactions with/to daylilies in any given instance. What is perfectly fine for one person may not be for another.)

In the previous installment of this series we looked at the traditional uses for daylily and some of the scientific research that is being done into the nutritional and medicinal uses of the daylily. While that look was in no way comprehensive, it is enough to demonstrate the uses of daylilies beyond the flower as a garden display, and it gives anyone interested some further reading suggestions to begin their own research.

In this segment I want to look at species and cultivars useful for those interested in growing daylilies for permaculture use, or who might want to locate base plants upon which to build a breeding program for any of those uses, whether solely for permaculture uses or even if just as a minor part of a daylily breeding program that encompasses other concerns with the flower as the major focus. I believe that the breeding and selection of traits valuable to nutraceutical concerns and permaculture applications do not have to be in conflict with breeding daylilies for floral gardens. For those interested the two can go hand-in-hand, as is true for any area of selection in a program. In spite of what we are sometimes told, it is possible to select for more than one trait at a time.

For those interested in daylilies in permaculture or food forests, the species and some of the hybrid cultivars may offer some excellent material to work with. However, if the garden has thrips and other insect pests, many daylilies from both groups will be problematic. In short, as the buds and open flowers are the major source of food and some of the medicinal uses, these insect pests damage your harvest, making them less palatable or destroying it partially or completely. The act of predation by these pests leave roughness, bumps, thorns and knots called enations on the bud, and in extreme cases, cause the buds to abort, sometimes even causing the scapes to wither entirely, loosing all the buds.  
Severe thrip damage to scapes on a highly susceptible daylily.

The outer shell of the bud is made up of the three outer petals (called sepals) of the flower. At the very least, there will be roughness on these and they can get very malformed due to this predation on susceptible plants. Buds that are ‘loose’, not tightly closed and perhaps partially open at the tip the day or two before the bud opens can allow these insects to get inside the flower and feed on the inner surface before the flower opens. It is my experience that while there are cultivars showing resistance to these insect predators, there seems to have been very little selection for this trait in the hybrid cultivars, probably because the majority of the wild species also show considerable susceptibility to these pests. However, there is resistance in some small number of hybrids and my own breeding work suggests that this resistance is heritable and can be intensified through crossing resistant individuals and selecting for the trait.

Spraying for pathogens is out of the question in true permaculture practice, and it is also out of the question in a resistance breeding program, so programs that do not spray, for whatever reason, are in the perfect situation to select for resistance to these pests. Many people reading this will be thinking that there are natural ways to control these pests, and there are. There are a number of insects that will predate these insect pests. However, in my garden, the heaviest thrip season falls across May and June, and with the advent of the heat of summer in early July I always see the activity of these pests go into decline. Unfortunately, the two major predators of thrips, aphids, etc, Praying Mantis and Lady Bugs, are not active at that time of year here. In my garden Praying Mantis and Lady Bugs are most active during the heat of summer when these pests have already gone into decline. This then defeats the use of natural predators. For me, the only path that has been very successful has been to select for resistant cultivars and then breed within them to select and increase resistance to these pest in the early and early mid-season flowering types. This is part of why I want to stress this so strongly. It is the only method that is really working for me, proving to be a real solution.

Here is a closeup of a seedling from my breeding program that shows extremely high thrip resistance. Note in this closeup that that the petal is literally swarming with thrips.

Here is a clump shot of the same seedling showing beautiful, clean flowers with no thrip damage and many buds on tall, well-branched scapes that also show no damage. This seedling also shows extremely high rust resistance. The parents are Ancient Elf x Solaris Symmetry (click the names to see information pages on these two cultivars). Ancient Elf shows extremely high rust resistance but thrip susceptibility, while Solaris Symmetry shows rust susceptibility but high thrip resistance. This seedling shows the desired combination of the resistance traits of both parents and demonstrates that such combinations can be achieved. In addition, this plant also shows the best foliage traits of both parents, being senescent with late emergence and beautiful, dark green coloring.

The same seedling above, here covered in seed pods. This seedling is very fertile, both ways, and so opens the door to a level of breeding that meets many of my personal goals. Note how deeply green and healthy the foliage is well after flowering has finished.

(Left) Ancient Elf and (Right) Solaris Symmetry
Parents of the above seedling.

While I love the many programs within the floral daylily community, I must say that going to modern hybrid daylilies for permaculture subjects is probably a loosing proposition, fraught with pitfalls. One runs the risk of wasting a lot of money on plants that require artificial care regimes, have little ability to compete in a more natural setting and tend to have little resistance to various pests and pathogens. As with pretty much everything else in the modern world, many floral daylily breeders practice ‘better living through chemistry’, and so have strayed for the natural, hardy roots of the species their domestic hybrid garden subjects descended from. While many modern hybrid daylily cultivars may be hardy and vigorous in the environments they are bred in, many are unlikely to be so in a permaculture or food forest settings. 

Mystic Butterfly
One of my favorite flowers I have ever grown. A truly stunning combination of floral traits - colors and forms.

However, when you pull away from the flower closeup, here is the reality of the plant in my garden. The foliage is always late-freeze damaged and the few scapes are always about ten to twelve inches tall with the flowers down in the foliage and seven to ten buds. Now, I am certain this plant flourishes in the garden in Florida where it was bred. I am certain in high-input care and/or warm-winter climates that it is a much better looking plant. I bet it would be stunning in a greenhouse. Yet, this is what it is in my garden. I am breeding with this plant in a salvage project though, because I absolutely love the flower and the plant does have some good traits here such as moderate rust resistance and it does not start growing at the first hint of warm weather. My goal is to bring flowers of this nature onto plants that are similar to the seedling pictures shown above. I want to stress again that this is not meant to be an incrimination of the breeder of this plant. I would not expect evergreen plants bred in Florida to show maximum performance in my cold-winter garden with average soil and low inputs. Such plants can't be selected for my conditions in vastly different conditions. That is simply reality and its not a bad place to reside.

Extreme inbreeding, over-focus on flower traits (and subsequently ignoring plant traits), the use of greenhouses and other artificial, chemically-focused, high-input growing environments and care regimen have caused the modern daylily cultivars to often be pale shadows of their wild ancestors in terms of survivability and ease of management. Never underestimate the ability of selection to do both good and bad things, depending on what is focused upon. While daylily hybridization and domestication has only been practiced in the west for a little over 100 year, that is ample time to bottleneck the gene pool and intensify deleterious traits through ignoring them in deference to the novelty of new visual looks in the flower.

Hemerocallis fulva Korean (Apps accession - 1984)
Growing in my hybridizing garden in two containers, which remain above ground all winter. Note the large, lush, healthy plants with many tall scapes. These two clumps are wildly pot bound, having been in those same pots for five years and I don't do anything to them, no fertilize or extra watering. The only thing I do is cut off the runners that try to come out of the holes in the bottoms of the pots on occasion. Containers are one excellent way to keep running fulva types in check. To read more about this fulva clone, visit my H. fulva Korean information page.

For the reasons cited in the last paragraph, and in combination with my own personal experience, I think there is actually little modern hybrid material that is suitable for use as a breeding base in developing daylily cultivars for permaculture settings. With that said though, I will also say that I firmly believe that any visual phenotype known in daylilies can be integrated into a program of breeding for permaculture use, IF the proper base plants are used to layer those visual traits upon AND proper attention is paid to a wide range of traits and proper selection pressure is applied to the seedlings. However, we do not have the wide range of data to tell us what effects, in terms of nutrients or chemical components, may be found in any given phenotype that has been heavily intensified through selective flower breeding. We don’t know, for instance, what effects the huge green throats that are so popular now many have. We don’t know what effect the selection toward blue-purple has intensified, or selection for bright, baby pink coloring may do to the nutraceutical traits. Personal and professional investigation is warranted to make further assessment. However, we know the species that are eaten and used medicinally in Asia, and so that is probably a good place to start.

Hemerocallis sempervirens
The latest blooming daylily in my garden, starting in Late August/September and going until frost in late October/November.

In my personal experience, the best base plants for breeding for permaculture uses are the clones of H. fulva (including the closely allied H. sempervirens) and H. citrina (including the closely allied H. vespertina).  Of the H. fulva clones, I find H. fulva ‘Korean’ and H. fulva ‘Hankow’ to be the most useful. The former shows less thrip resistance, but can produce resistant seedlings, especially when bred with other plants showing thrip resistance. The later is late blooming and so tends to miss the thrip season in my garden. It can throw both late blooming seedlings, and when bred with earlier blooming cultivars with thrip resistance, can also produce earlier-flowering resistant seedlings. Other fulva clones are probably useful, and I am currently testing out several other forms, but ‘Korean’ and ‘Hankow’ seem to be exceptional for breeding in my program. 

Of all the fulva clones, ‘Europa’, ‘Kwanzo’ and ‘Flore-pleno’ seem to show the least utility in any type of breeding, as they are near-sterile triploids. My experience is that I can’t produce enough seedlings from any of them to actually apply the type of selection I wish to make. Others may find them more useful. While ‘Korean’ and ‘Hankow’ are almost certainly triploids as well, they are quite fertile with both diploids and tetraploids in my own personal experience (and that of others) and so can be used in programs at either diploid or tetraploid levels. H. sempervirens may be its own species or it may be some form of fulva, but whichever it actually is, it is a good breeder and is very late flowering, even later than ‘Hankow’. Whether it has thrip resistance or not, I don’t know, but I have never seen problems with insect predation on it. This may be because of the late flower. It is probably a diploid, and I have produced numerous seedlings with it when crossed with pollen of diploid cultivars. I haven’t yet used it with tetraploid pollen, so can’t yet speak to that. Amongst its seedlings, some are earlier flowering and they also have not shown much insect predation, but that may be because they are still flowering too late to have heavy exposure to the problem. I do think H. sempervirens is useful, especially for breeding late flowering plants, and it is heavily branched with many, many buds and a long flower season that is later than any of the other species I know of.
A clump shot of H. vespertina showing the extremely tall and branched scapes and the large plant. A late bloomer, typically starting in August and going for about six weeks.

A closeup of the flower of H. vespertina on an immature plant showing less branches, buds and lesser height of scape than is to be expected on a mature clump. The flower is typical of the mature plant.

Amongst the yellow types, I have found all the clones of H. citrina I have grown to have high thrip susceptibility. H. vespertina, which may be a form of citrina or may be its own species, is far superior to all the citrina clones I have grow, perhaps because it blooms quite late in the season. It is very tall and heavily branched with many buds. Seedlings I have produced from it have also tended to show resistance to insect predation, however, this again many be due to their flower in July and August. The yellow species are thought to be diploid. While one might get an occasional set from tetraploid pollen, this would be a very rare event. I haven’t yet tried H. vespertina with tetraploid pollen, and I spent an entire summer putting tetraploid pollen on H. citrina clumps to get two pods with seventeen seeds. While one might persevere and create plants with chromosome counts above the diploid level using the yellow species, generally they are easiest to use at the diploid level.

Notify Ground Crew early morning 8 AM

Notify Ground Crew afternoon 5 PM

Notify Ground Crew at sunset 9 PM

Notify Ground Crew at night 11 PM

For yellow near-species/species-like hybrid cultivars my favorite cultivar as a base plant is Notify Ground Crew (click name to see information page about this cultivar). It is a large plant with tall (6’) scapes, well branched and with many buds. It shows much greater thrip resistance than the citrina clones I have grown, and it descends from a tetraploid conversion of H. citrina made by Orville Fay (via pod parent Tetrina’s Daughter) on one side and tet. Purity on the other as pollen parent, which is also close to species. Notify Ground Crew is a great breeder and I have produced many seedlings from it that also show strong thrip resistance. Another useful hybrid cultivar, descending from two converted species-like yellow parents, is Ancient Elf, though it has fairly high thrip susceptibility in my garden. Crossed with plants showing high thrip resistance, it can produce much more resistant offspring and it has many other good plant traits including very high rust resistance.
In addition to thrip and other insect resistance, most of the species-clones and cultivars I discussed above tend to have moderately high to very high rust resistance. This may be useful for some programs where that is also a concern, such as in the deep south or anywhere that daylily rust can survive and there is no winter freeze to kill the rust fungus. 

A daylily plant showing heavy daylily rust infection. Such a plant is susceptible to the rust fungus. Some plants show resistance to this fungus. For more information on cultivars resistant to daylily rust and breeding for rust resistance, click here.

Another point about all those mentioned above is that they can be grown throughout a wide range of climates. Daylily cultivars are registered as ‘dormant’, ‘semi-evergreen’ and ‘evergreen’. These categories are very loose and are not botanical descriptions, but are hobby shorthand. While some think these are hard-and-fast categories there are no true, binding botanical rules as to what gets registered how, and so they are not reliable guides. However, plants having leaves that die in fall and then form a resting bud may require some cold and many of these fail in southern gardens. Some plants retain some amount of foliage, green, above ground throughout the winter regardless of climate and only vanishing to or near the ground when they are frozen off. Many of these are hardy in cold climates, but others are tender and are harmed or even killed by freezing conditions. 

This picture shows a clump of Ancient Elf in the winter. Note that you can see the name tag, but you don't see any plant. This demonstrates the full senescence of the foliage. The resting bud is below the dead leaves and just at or under the ground level and will begin to emerge when spring commences, showing moderately late emergence allowing it to skip many freeze/thaw cycles. Daylilies of this nature may fail in areas without sufficient winter cold.

The fulva clones and citrina clones are adaptable. In cold climates they tend to go into dormancy, with only a tiny amount of plant above ground or only a resting bud, allowing them to thrive in very cold climates. In warm climates they tend to be ‘semi-evergreen’ or ‘evergreen’, and this also allows them to thrive in even very warm climates. For this reason, these species are recommendations I can make for almost any climate. Notify Ground Crew behaves much as the citrina and fulva types, though it is registered as ‘dormant’. Of all those I mentioned above, H. sempervirens is the most evergreen (sempervirens literally means ‘evergreen’) and while it survives in the far north, it doesn’t thrive, and Ancient Elf is the most dormant, with foliage that dies back in fall forming a resting bud for a period of time. It may not thrive in the deep south. While the species clones and hybrid cultivars I have mentioned here have strong applications for permaculture settings, they also have excellent applications in floral breeding programs, opening the gene pool of inbred and bottlenecked lines and possibly bringing genes not found in the modern hybrids into those lines.

Here is a picture of an evergreen type in my garden in winter. Some evergreen daylilies are hardy in my garden, while others are not and are referred to as 'tender'. These daylilies tend to attempt growth at each warm period throughout the winter and early spring and can sustain repeated damage that can reduce their performance. Such daylilies prosper in warm-winter climates such as the deep south of the US, and while some are perfectly hardy in the cold-winter climates, some are tender and not able to thrive or survive in repeated freezes. Ironically, some of these evergreen types do better in the far north than they do in my zone 6/7 garden, as some northern climates provide an insulating blanket of snow throughout the winter, protecting these plants from beginning growth repeatedly before spring has fully arrived. My area never has the long-lasting, insulating blanket of snow.

In part three we will look at going beyond the base plants and developing programs targeted at specific phenotypes.